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Thanks are due to all people involved in the field work over the years: César Cajigal, Alejandro Isla, Fredrik Nordwall, Felipe Reyes-Gavilán, Antón Fernández, Ataulfo Martínez, F. Cimentada. The staff of Picos de Europa National Park kindly helped with site selection and administrative tasks. Funds were provided by grants FAIR CT95-0009 to F. Braña and FAIR CT97-3498 and MCYT-AGL2000-3181-CE to A. G. N. We thank Lucie Buttay for helping with data management and Daniel Oro, Marc Mangel and one anonymous referee for their useful comments on earlier versions of the manuscript. A. F.-C. was supported by a FPU grant from the Spanish Ministry of Education (ref. AP2008-04476). We also thank Stuart Larsen for revising the English